Compilation of key molecular reporter lines published in literature to study cell and developmental processes in
Mosses are plants that belong to the Bryophytes, a cosmopolitan sister group of vascular plants with the last common ancestor between 400 and 500 million years ago [1, 2]. As a mostly avascular lineage, Bryophytes, that include mosses, liverworts and hornworts, thrive in mostly moist niches near the surface and stay compact (<10 cm), with some neovascularised exceptions that grow up to 65 cm [3, 4, 5]. Their cosmopolitan distribution in a variety of biotopes including moist and arid environments, can be explained by unique adaptations like drought, freezing and salinity tolerance [4, 6, 7]. Mosses’ life cycle is dominantly gametophytic (the photosynthetic and growing phase is haploid), and the size and architecture of their organs is smaller and simpler than that of vascular plants, with leaf-like structures (phyllids) and sexual organs (antheridia and archegonia) of often only one cell of thickness, stem-like structures of circa ten cells and spore-bearing containers (sporangia) of single-cell spores [4, 8, 9, 10].
This miniaturised body renders mosses accessible systems for the study and dissection of cell and molecular aspects of plant biology that require close monitoring in time and space [11]. Such studies greatly benefit of the accessibility to single cells in a multicellular context. For instance, asymmetric and directional cell divisions are key life developmental tools to build an organism, but we lack understanding on how these processes are exactly controlled and regulated [12]. Importantly, these developmental drivers are shared between mosses and vascular plants, and to some extent with animals, and associated gene functions seem to be highly conserved despite the long period of independent evolution [13].
In the last two decades, mosses have gained high interest in plant research, with
Hereby, we present how mosses, thanks to their simplified body plan and genetic networks in development, and with special focus in
As most land plants, mosses have alternating generations between the haploid gametophyte and diploid sporophytes. However, unlike vascular plants, mosses spend most of their life cycle in the gametophytic stage, in which most of the organism asexual development, including photosynthesis and growth, occurs. The sexual organs eventually develop at this stage to give rise to the embryo after fertilisation, that produces the sporophyte over the gametophyte. This fruiting stage is diploid until haploidisation in spore formation takes place [11, 14].
Starting from a spore, the first developmental stage of the moss is the chloronema, a chloroplast-rich and single cell-wide filamentous tissue that serves for initial colony expansion, early photosynthesis and nutrient absorption. The cells are slightly elongated (~80 μm long), and the intercellular cell walls are oriented perpendicular to the growth direction [15]. This filament eventually transitions to caulonema, a quick-growing filamentous cell type that have underdeveloped chloroplasts at early stage, with longer and narrower cells (~250–300 μm long) that grow twice as fast, and with oblique intercellular cell walls [15, 16, 17]. This tissue has exploratory purposes and is favoured in stressful, light-poor, and nutrient-poor conditions, possibly with the aim of finding more suitable conditions [18, 19]. Caulonema can transition again to (secondary) chloronema [17]. The filamentous tissues are collectively referred to as protonemata and can laterally grow and divide to branch as new filaments. The protonemata grow in a mat-like fashion that shapes the two-dimensional (2D) developmental stage of mosses where the growth is confined in a plane of few millimetres of thickness.
Sometimes, the lateral cell outgrowth (the cell initial) gives rise to a bud cell instead of a branch cell, which is the beginning of gametophore development and the transition to three-dimensional (3D) growth [20]. The identity of the cell initial can be predicted by the division plane angle, implying that identity is determined before division (Figure 1). Currently, the list of known genes involved in the path selection and division plane orientation is growing, but the early determinants of bud formation and branching remain unknown [18, 21]. The bud grows by well-defined asymmetrical and oriented divisions to form the gametophore, the leafy shoot-like plantlet of mosses that ultimately bears gamete-producing organs.
Scheme of morphology and location of different developmental processes.
The bud basal cell gives rise to a new type of filamentous tissue, the rhizoids, with pigmented, caulonema-like morphology. They function as anchorage to the ground to stabilise the up-growing gametophore and contribute to nutrient and water uptake, similar to roots and root hairs of seed plants, but with the tissue complexity of root hairs [22, 23]. The bud apical cells divide in precise directions to give rise to oriented phyllid initial cells with a particular phyllotactic pattern (i.e. lateral organ organisation around the shoot; e.g. spiral) to develop the gametophore.
The apical cells eventually arrest their proliferation, or terminally give rise to sexual organs (firstly antheridia, and later archegonia) under autumnal/spring conditions: short day (8 h), low light (20 μmol/m2/s) and low temperature (15°C) [24, 25]. Despite the asynchronous development of male and female gametangia, this moss is self-fertilising, and thus tends to genetically self-isolate [26]. Flagella-driven spermatozoids (male gametes) move towards the archegonial venter in liquid water and fertilise the egg cell to give rise to a diploid zygote. The zygote will subsequently develop, via an embryonic stage with a new 2D-to-3D transition, into the sporophyte, that consists of the foot (the interface with the gametophyte) and a short stalk (seta) with a terminal capsule. In the capsule, meiosis gives rise to up to few thousands of haploid spores [8, 26].
The first documented ecotype, known as ‘Gransden’ (United Kingdom, 1962), has reduced rates of sporophyte formation, probably due to long asexual propagation in laboratories [27]. In many laboratory lineages, it has become self-sterile, rendering it unattractive for studies dependant on sexual reproduction. On the contrary, the more recently isolated ecotypes ‘Villersexel’ (France, 2003) and ‘Reute’ (Germany, 2006) have 15 times more sporophytes (77% of total gametophores), indicating a high fertility rate [26]. Despite these differences, all ecotypes can be propagated asexually in identical conditions from any tissue thanks to the high regeneration rate of this moss, through which explants will redifferentiate into chloronema and initiate the life cycle from there [28, 29].
Several essential developmental processes are shared between all land plants, including angiosperms and bryophytes. Strikingly, in mosses many shared processes take place with a simplified set of genes and sometimes in single cells. Therefore, the underlying genetic regulatory networks of development are easier to study. In this section, we highlight some of these processes and their unique ease of study in the moss
The ability to structure organs in three dimensions (3D) was an essential feature for water-to-land transition. Many aquatic plants develop in a homogeneous environment, whereas land plants faced a highly distinct environment at ground surface level (a plane) and at the air/soil axis (perpendicular to this plane). This required land plants to develop specific tissues to efficiently grow in each dimension and cope with new challenges [20].
From a physiological perspective, the transition from 2D-to-3D growth in mosses consists in the development of complex structures such as the gametophore that grows out of the surface plane where protonema thrive, exhibiting negative gravitropism and positive phototropism [30, 31, 32]. The development of the gametophore relies on the ability of the moss to define different organisation in each dimension of space. The basal units from which this spatial assembly takes place are ultimately single cell primordia, from which organs emanate [33]. To this end, a cell must spatially sense intrinsic and extrinsic signals and transduce them to subcellular structures that pave the way to division plane positioning and subsequent asymmetric and oriented cell division. However, the molecular basis of spatial sensing and its transduction to organisation, action and maintenance of the division machinery has not been fully elucidated yet [12].
Cellularly, the 2D-to-3D transition in a moss occurs during bud formation (Figure 1). When a cell initial is formed in protonemata, in 5% of the cases it has a bud initial cell identity instead of a branch initial identity. Each bud initial swells and undergoes divisions oriented in all three axes of space [34]. This transition is supported and maintained by a different genetic and molecular machinery than that of protonemal development, and is the molecular basis of 3D growth. These proteins are specifically present from the first cell of the bud and onwards (e.g. DEK1, NOG1), and remains active in subsequent proliferating organs (e.g. shoot apical meristem, phyllids, gametangia) [13, 34, 35, 36]. In general, gradients or local clusters of proteins, peptides, nucleic acids, and hormones can be signals, sensors and/or actuators in developmental processes, and what is upstream of the cascade of 2D-to-3D transition remains a mystery. Some actors that are already on the radar include oscillations of auxin and cytokinin concentrations, ROP and SOSEKI proteins and CLE/CLV peptides-receptors, and they have been pinned down to different moments of the process and at different locations [12, 37, 38]. However, how do they coordinate their activity remains elusive and an active area of study.
In vascular plants, the 2D-to-3D transition occurs only once in their lifecycle, during embryogenesis, where orthologs of the essential moss 3D machinery genes are also expressed [34, 35]. This event is confined in the endosperm during embryogenesis and its observation in seed plants requires seed and ovule microdissection, which makes
These features provide a privileged seat for
The protagonist of 2D-to-3D growth transition is the formation of the shoot apical meristem (SAM), a region at the apical growth side responsible for the continuous formation of the aerial organs of the plant, including leaves and reproductive organs [40, 41]. The histology of the SAM in angiosperms describes a central zone with stem cells that self-renew, divide and radially differentiate into peripheral cells that determine organ initiation at specific locations (e.g. by placement and establishment of new leaf primordia) [42]. While angiosperms present multicellular stem cell centre SAMs during their sporophytic phase, bryophytes present an unicellular structure, the shoot apical cell (SAC), both in their gametophytic and sporophytic phase (Figure 1) [43]. Despite the differences, both SAM/SAC share a common organisation, with stem cell(s) at the centre, surrounded by regularly differentiating tissue [44].
The mechanisms that establish and maintain these pluripotent stem cell(s) in the SAM/SAC is unknown. It involves spatial sensing, cell-to-cell communication, and asymmetric and oriented cell divisions, which render mosses attractive systems for their accessibility.
The developmental origin of the sporophytic SAC in
The genetic make-up in both taxa has shown to rely on auxin response through AUXIN RESPONSE FACTORS (ARFs), cytokinin signalling by ARABIDOPSIS RESPONSE REGULATORS (ARRs), CHASE domain-containing histidine kinases (CHKs) and CYTOKININ OXIDASE/DEHYDROGENASE (CKX), local coordination through several transcription factors families like CLAVATAs (CLVs), CUP-SHAPED COTELYDONS (CUC), LATERAL ORGAN BOUNDARY (LOB) and signalling peptides like CLAVATA3/EMBRYO SURROUNDING REGIONRELATED (CLE) and chromatin modification by Polycomb Repressive Complex 1 and 2 (PRC1,2), among others [48]. Although many key factors have conserved roles in SAM formation and maintenance in seed plants and mosses, some important factors in angiosperms, like the key regulator of stem cell maintenance WUSCHEL, are not found in
The most noticeable outcome of the shoot apical meristem/cell (SAM/SAC) activity is the organised and oriented initiation of leaf primordia along the stem, which leads to a unique geometric pattern of leaves and shoot branches named
However, the pattern arises from essentially different SAMs/SACs: in angiosperms, phyllotaxis derives from a multicellular system with well-reported oscillating auxin peaks around the SAM growth axis, whereas
The limited understanding of the origin and underlying molecular mechanisms of this rotating pattern and derived phyllotactic pattern is largely confined to the sporophyte of the evolutionarily recent group of flowering plants (angiosperms). The available transcriptomic data of bud and tip cells and gametophores (or 3D shoots) may provide more insight in the transition from uniplanar to triplanar meristematic growth in moss [48].
Aligned with the phenotypic similarities of moss and angiosperm phyllotactic patterns, several factors known from Arabidopsis have also been found in mosses, including receptor signalling genes involved in shoot meristem size and patterning, hormone biosynthesis genes, transcription factors that control cell-specific mechanism of developmental pattering, chromatin remodelling complexes and cell cycle [48]. Many of these factors are essentially executive and likely controlled by some spatial sensing machinery. Comparing them with new contributors or absent members in the minimal regulatory network of mosses may help unravelling the fundamental elements that trigger the orientation-specification machinery that greatly impacts plant architecture in all land plants, including relevant crops.
Most described processes in this chapter require cell identity acquisition and maintenance. In certain circumstances (e.g. wounding), differentiated plant cells can reprogram to become new stem cells, divide and redifferentiate for organ
In most tissue cultures of other plant species, exogenous hormones (e.g. auxin and cytokinin) are required to induce callus formation and plant regeneration [60]. However, in moss, cells are capable of regenerating from protoplasts or excised phyllids into new protonema filaments in the absence of exogenous hormones (Figure 1) [61]. This implies that the whole regeneration toolkit is present in mosses and can be endogenously activated on demand, which makes them different from other taxa (e.g. angiosperms) [62].
When a phyllid is excised, cells neighbouring the cutting edge can reprogram from somatic cells to protonemal stem cells, which can then start a new life cycle [14]. This regeneration process is easy to study in mosses for several reasons: firstly, cell identity conversion can be easily tracked with protonema stem cell reporters [29]; secondly, aside of the simplicity of
The mechanistic studies of the cell fate acquisition can benefit from this simple cell type conversion in comparison to other model systems used in cell reprogramming investigation (e.g. regenerative callus or Arabidopsis roots that consist of multiple cell types that possess different tissue identity) for its minimality and event frequency [65, 66]. In angiosperms, regeneration is often reduced to localised stem cell pools (e.g. the base of leaves), takes longer to establish, and it is multicellular and asynchronous at the explant level [58, 67].
Previous studies have taken advantage of the abovementioned features to investigate gene expression profile during phyllid cell reprogramming, which revealed that genes involved in stress, proteolysis, and hormone signalling pathways are induced from 6 to 24 h after cutting [64]. Some genes have been demonstrated to play essential roles in moss leaf reprogramming, including
One interesting feature of moss regeneration is inhibition of neighbouring cells. The necessary cell–cell (apoplastic e.g. Ca2+-mediated) or cell-to-cell (plasmodesmata-mediated) communication makes regeneration an attractive developmental process to study this cell crosstalk [71, 72, 73]. The phytohormone ABA is a key responsible of the dynamic regulation of the permeability of plasmodesmata in response to changing environments, such as wounding. Control in plasmodesmata pore size can influence the signalling molecules that can pass through or can be blocked in particular cells, which can have a direct effect in development of the processes mentioned until now [74, 75].
The signalling pathways and functions of plant hormones are substantially conserved in
Three plant hormones—auxin, cytokinin and strigolactone—have shown to regulate shoot branching patterns (phyllotaxis) and activation in angiosperms. Auxin moves down the main shoot of angiosperms to inhibit branch development, while cytokinin promotes branching. In addition to branching, auxin is the key molecule in the control of plant growth and development, and promotes organ differentiation [76, 77]. Exogenous application of auxin or its inhibitors results in irregular cell shapes and inhibit lateral organ formation, for instance in shoot apical meristem (SAM) maintenance. The understanding of hormone regulation and signalling in angiosperms progresses slowly due to tissue and gene network complexity. In
Cytokinins and strigolactones also influence plant architecture, both in angiosperms and mosses. The levels of cytokinin are high and precisely distributed in the central stem cell region of SAM in angiosperms to maintain stemcellness [84]. In the root apical meristem, auxin and cytokinin act antagonistically in meristem size control, but its levels, distribution and interaction in
As shown, many processes that define plant architecture are regulated in similar ways both in angiosperms and mosses. However, mosses offer a reduced gene network and regulation complexity that facilitates the analysis of hormone functions in the related developmental processes. Furthermore, subcellular and tissue-level transport and distribution of hormones can be best visualised in their simple plant bodies. In such plant models, new hormone functions will prove to be easier to study and translate to agronomically relevant plants.
Many valuable online resources with information on protocols, stocks, tools and genetic information have been exhaustively compiled elsewhere [11]. Hereby, we provide some additional information and summary of the essentials of research in
The small size and simple architecture of moss organs allows detailed microscopic visualisation easy to accomplish in almost all tissues. The strings of cells in protonema and their branching is trivial to closely visualise, and the transition to gametophores can be well tracked until the stem-like centre becomes slightly thicker than a dozen of cells and grows out of the plane. From it, the leaf-like structures (phyllids) have only one cell of thickness except in the midrib and can be tore apart for up-close visualisation. The terminal sexual organs (antheridia and archegonia) have a 3D structure that is easy to fully dissect or directly visualise due to their monolayered sack structures. The subsequently developed spore-bearing containers (sporangia) are full of single-cell spores [4].
This miniaturised body renders mosses accessible systems for the study and dissection of cell and molecular biology that require close monitoring in time and space. Such studies greatly benefit of accessibility to single cells in their context for observation of subcellular responses
In
Visualised | Fusion/Target | Purpose | Reference |
---|---|---|---|
Nucleus | NLS4-GFP-GUS | Nuclear localisation | [21] |
Endomembrane system | Endoplasmic reticulum | [11] | |
α-1,2-mannosidase | Golgi apparatus imaging | [88] | |
Targeting signal type 1 (SKL) | Peroxisome imaging | [88] | |
Mitochondria | Cytochrome c oxidase | Mitochondria imaging | [88] |
Cytoskeleton | LifeAct | Actin cytoskeleton | [89] |
Tubulin α | Microtubule cytoskeleton | [90] | |
MAP65 | Antiparallel Microtubule-microtubule contacts | [91] | |
Kinesins | [90] | ||
Plasma membrane | SNAP-TM-mCherry | Membrane tracking | [21] |
Auxin | pDR5v2:GFP-GUS | Aux. induced fluor. | [92] |
GH3:GFP-GUS | Aux. induced fluor. | ||
R2D2 | Ratiometric induction | ||
Protonema | pRM09:NLS4-GFP-GUS | Protonema identity reporting | [29] |
pRM55:NLS4-GFP-GUS | Protonema identity reporting |
Compilation of key molecular reporter lines published in literature to study cell and developmental processes in
Despite the advantageous physiological features of
Microfluidic devices are transparent and flexible structures commonly produced using the biocompatible and air-permeable polydimethylsiloxane (PDMS) polymer. In biology, they have been used for study and imaging of cell and tissue development
Until now,
Some groups carried out forward genetic screening by X-ray or chemical mutagens that generated mutants with hormone resistance or abnormal tropic responses [85, 98]. However, due to the lack of genomic information, the disrupted genes that caused the phenotypes were never identified.
Recently, the completion of genome sequencing and the establishment of its genetic mapping tools removed the obstacles in the forward genetic screening of
With this genetic mapping resource, in the past few years, researchers started to perform forward genetic screening by treating protoplast with UV light. In such screenings, mutants with phenotypic defects were successfully obtained and the causal lesions were identified by outcrossing and whole genome sequencing. Notably, essential genes that are crucial for growth may not be identified due to the lethality of their knock-outs; therefore, conditional screening was performed to overcome this problem. After the UV light treatment, plants were cultured under different temperatures and in such conditions, plants showed growth defect only in high temperature were selected as a temperature-sensitive mutant [100].
Another screening aimed to discover genes that are essential for the 2D-to-3D transition is also limited by developmental defects, given that mutants in this process cannot produce gametophores necessary for sexual crossing. To overcome this problem, instead of crossing, researchers generated somatic hybrids between Villersexel mutants and Gransden wildtype, which produced diploid sporophytes [34]. Spores released from this hybrid sporophyte exhibited consistent phenotypic segregation ratio with meiosis. Mutant plants generated from these diploid spores were sequenced and genomically mapped to achieve the identification of new crucial genes for moss 2D-to-3D transition (e.g. NO GAMETOPHORES 1 and 2, or NOG1, NOG2).
In addition to UV light, tobacco Tnt1 retrotransposon was used to produce insertional mutations in genic and GC-rich regions [101]. Both PEG- or Agrobacterium-mediated transformations were applied and successfully produced mutants.
The integration of DNA constructs (including promoter, gene of interest and selection cassettes) necessary to produce transformants with stable expression and non-disrupting phenotypes requires targeting of neutral loci that do not intrinsically produce a phenotype when disrupted, often due to gene redundancy. In Table 2, there are several standard neutral loci indicated which reportedly showed no visible phenotypes or morphological defects when it is replaced by an entire gene expression cassette.
Locus | Vector | Purpose | Reference |
---|---|---|---|
PIG1 locus | pGX8 pGG626 | XVE inducible overexpression XVE inducible RNAi expression | [102, 103] |
Pp108 locus | pUGGi | Constitutive RNAi expression | [104] |
Pp108 locus | pTH-Ubi-Gate | Constitutive expression by the maize ubiquitin promoter | [105] |
Redundant copy of the ARPC2 gene | pTK-Ubi-Gate | ||
Redundant copy of the ARPC3 gene | pTZ-Ubi-Gate | ||
PTA1 locus | pT10G | Overexpression by EF1α promoter | [106] |
BS213 locus | pMJ1 | [107] |
A compilation of vectors designed to target proven neutral loci in
Currently there are several vector sets released to specifically target neutral loci, that contain their flanking regions homologous to parts of the locus at start and end of the vector. Cloning the gene of interest in between readily allows replacement of the targeted locus with the entire cassette via homologous recombination (see section
In
The game-changing CRISPR/Cas9 method has proven an efficient and effective tool in
In Nogué’s lab, a co-delivery method was developed where each element of the system (Cas9, sgRNA and selection cassettes) was present in a separate plasmid. In this method, Cas9 expression is driven by an actin promoter and ready to use as is. The selection strategy can be chosen freely due to the lack of integration, minding the presence of resistance in the background lines. This method is also suitable for multiple mutations in different genes at once, given that more than one sgRNA plasmid can be simultaneously delivered in one transformation with still sufficiently high efficiencies of transformation [110].
In Bezanilla’s lab, a whole set of gateway destination vectors for CRISPR/Cas9 system was developed. The strategy was to design a vector set to finally put all the three essential components (Cas9, sgRNA and selection cassettes) in a single expression plasmid. In both protocols, the sgRNA can be designed and optimised using the online design tool CRISPOR V1 against
To increase the accuracy of mutations, the CRISPR/Cas9 system is applied with a homology-directed repair (HDR), which allows for seamless knock-in or point mutation in desired sites. The template DNA can be a donor plasmid that harbours homologous fragments or oligodeoxynucleotides (ODNs) [111, 113]. By co-transforming the plasmid or ODNs together with CRISPR/Cas9 and sgRNA vectors, both methods show high accuracy to generate a desired point mutation or scarless insertion with a fluorescence tag at any suitable location of the gene.
Given that moss possesses a relatively big gene family, arguably due to its double genome duplication, the employment of gene deletion strategies might be inefficient to investigate gene functions due to the high redundancy rate (e.g. there are four ROP genes with highly homologous or identical sequences) [38]. For this, RNA interference (RNAi) strategies offer an alternative to overcome this problem, and the procedure has been well-established.
To generate an RNAi construct for a gene of interest, a DNA sequence of 300 to 1000 bp is subcloned in a destination vector. After standard PEG-mediated transformation, the silencing effect can be detected after 24 h and last up to 3 weeks [104]. To avoid lethal effects when constitutively expressing interfering RNA, an oestradiol-inducible RNAi system is available [102]. Coupling RNAi silencing activation with fluorescent reporters facilitates screenings of loss of function phenotypes [104].
Standard transformation protocols have been applicable to
Strategy | Highlights | Reference |
---|---|---|
PEG/Mannitol | One to two round selection, 10% of transformants, 3–4 weeks for stable transformants | [114, 115, 116, 117] |
Four-round selection, 100% positive transformants, 12–16 weeks | [118] | |
Particle bombardment | Easy to conduct; less used. Transient and stable DNA integration. | [119] |
Summary table of the classical and current transformation techniques and reference protocols for application.
Due to their accessibility, tractability and close yet independent phylogenetic position, the interest in Bryophytes has increased dramatically in the last decade [1]. Beyond
The fire moss
The Hypnales (Bryopsida) are the biggest and most diverse order of mosses with varied morphology, and mostly exhibit pleurocarpous (i.e. non-erect) growth fashion. It includes
Remarkably, the common aquatic moss
The genus
The moss class Polytrichopsida is the second biggest (~200 species) after Bryopsida (~11500 species), and its species have unique morphological characteristics that make them uniquely interesting for developmental biology [5]. Despite mosses being regarded as avascular plants, some exhibit hydroids and leptoids, a functionally analogous tissue to tracheids and sieve elements of vascular plants that slightly differs morphologically and developmentally. Polytrichopsida has several genera with such structures, in some cases underdeveloped and, in others, completely functional, like in
Despite its potential to provide valuable insight, genetic tools have barely been developed for this taxon, but full mitochondrial and plastid genomes have recently been published for
The desert moss
The heavy metal-tolerant moss
We have shown how mosses are an increasingly relevant model group to plant developmental biology due to their distinct accessibility at physiological and molecular level. Their evolutionary distance with agronomically relevant plants does not diminish their potential to help to understand fundamental questions of development that remain unsolved, given that most essential regulatory networks are conserved. This has been shown in the hormonal regulation of branching and shoot and root development, regeneration, the establishment of shoot apical cells and the genetic make-up in 2D-to-3D transition. The utility of mosses has convinced the scientific community to the point of promoting the sequencing of eight new species to explore other physiological processes in the last few years. We have also shown how
The authors would like to thank Dr. Jeroen de Keijzer and Dr. Tijs Ketelaar for their thoughtful and detailed review of the manuscript. Also, the funding agencies Technology, Knowledge and Innovation, division Horticulture and Propagating Material (TKI T&U) and the Dutch Research Council (NWO) (reference number: TKILWV20.390) for funding JFC and the ERC grant to Prof. J. Friml (reference number: PR1023ERC02) for funding HT. The authors would like to sincerely apologise for the literature not cited that may be relevant for this chapter and is not present due to space constraints.
Mosses are plants that belong to the Bryophytes, a cosmopolitan sister group of vascular plants with the last common ancestor between 400 and 500 million years ago [1, 2]. As a mostly avascular lineage, Bryophytes, that include mosses, liverworts and hornworts, thrive in mostly moist niches near the surface and stay compact (<10 cm), with some neovascularised exceptions that grow up to 65 cm [3, 4, 5]. Their cosmopolitan distribution in a variety of biotopes including moist and arid environments, can be explained by unique adaptations like drought, freezing and salinity tolerance [4, 6, 7]. Mosses’ life cycle is dominantly gametophytic (the photosynthetic and growing phase is haploid), and the size and architecture of their organs is smaller and simpler than that of vascular plants, with leaf-like structures (phyllids) and sexual organs (antheridia and archegonia) of often only one cell of thickness, stem-like structures of circa ten cells and spore-bearing containers (sporangia) of single-cell spores [4, 8, 9, 10].
This miniaturised body renders mosses accessible systems for the study and dissection of cell and molecular aspects of plant biology that require close monitoring in time and space [11]. Such studies greatly benefit of the accessibility to single cells in a multicellular context. For instance, asymmetric and directional cell divisions are key life developmental tools to build an organism, but we lack understanding on how these processes are exactly controlled and regulated [12]. Importantly, these developmental drivers are shared between mosses and vascular plants, and to some extent with animals, and associated gene functions seem to be highly conserved despite the long period of independent evolution [13].
In the last two decades, mosses have gained high interest in plant research, with
Hereby, we present how mosses, thanks to their simplified body plan and genetic networks in development, and with special focus in
As most land plants, mosses have alternating generations between the haploid gametophyte and diploid sporophytes. However, unlike vascular plants, mosses spend most of their life cycle in the gametophytic stage, in which most of the organism asexual development, including photosynthesis and growth, occurs. The sexual organs eventually develop at this stage to give rise to the embryo after fertilisation, that produces the sporophyte over the gametophyte. This fruiting stage is diploid until haploidisation in spore formation takes place [11, 14].
Starting from a spore, the first developmental stage of the moss is the chloronema, a chloroplast-rich and single cell-wide filamentous tissue that serves for initial colony expansion, early photosynthesis and nutrient absorption. The cells are slightly elongated (~80 μm long), and the intercellular cell walls are oriented perpendicular to the growth direction [15]. This filament eventually transitions to caulonema, a quick-growing filamentous cell type that have underdeveloped chloroplasts at early stage, with longer and narrower cells (~250–300 μm long) that grow twice as fast, and with oblique intercellular cell walls [15, 16, 17]. This tissue has exploratory purposes and is favoured in stressful, light-poor, and nutrient-poor conditions, possibly with the aim of finding more suitable conditions [18, 19]. Caulonema can transition again to (secondary) chloronema [17]. The filamentous tissues are collectively referred to as protonemata and can laterally grow and divide to branch as new filaments. The protonemata grow in a mat-like fashion that shapes the two-dimensional (2D) developmental stage of mosses where the growth is confined in a plane of few millimetres of thickness.
Sometimes, the lateral cell outgrowth (the cell initial) gives rise to a bud cell instead of a branch cell, which is the beginning of gametophore development and the transition to three-dimensional (3D) growth [20]. The identity of the cell initial can be predicted by the division plane angle, implying that identity is determined before division (Figure 1). Currently, the list of known genes involved in the path selection and division plane orientation is growing, but the early determinants of bud formation and branching remain unknown [18, 21]. The bud grows by well-defined asymmetrical and oriented divisions to form the gametophore, the leafy shoot-like plantlet of mosses that ultimately bears gamete-producing organs.
Scheme of morphology and location of different developmental processes.
The bud basal cell gives rise to a new type of filamentous tissue, the rhizoids, with pigmented, caulonema-like morphology. They function as anchorage to the ground to stabilise the up-growing gametophore and contribute to nutrient and water uptake, similar to roots and root hairs of seed plants, but with the tissue complexity of root hairs [22, 23]. The bud apical cells divide in precise directions to give rise to oriented phyllid initial cells with a particular phyllotactic pattern (i.e. lateral organ organisation around the shoot; e.g. spiral) to develop the gametophore.
The apical cells eventually arrest their proliferation, or terminally give rise to sexual organs (firstly antheridia, and later archegonia) under autumnal/spring conditions: short day (8 h), low light (20 μmol/m2/s) and low temperature (15°C) [24, 25]. Despite the asynchronous development of male and female gametangia, this moss is self-fertilising, and thus tends to genetically self-isolate [26]. Flagella-driven spermatozoids (male gametes) move towards the archegonial venter in liquid water and fertilise the egg cell to give rise to a diploid zygote. The zygote will subsequently develop, via an embryonic stage with a new 2D-to-3D transition, into the sporophyte, that consists of the foot (the interface with the gametophyte) and a short stalk (seta) with a terminal capsule. In the capsule, meiosis gives rise to up to few thousands of haploid spores [8, 26].
The first documented ecotype, known as ‘Gransden’ (United Kingdom, 1962), has reduced rates of sporophyte formation, probably due to long asexual propagation in laboratories [27]. In many laboratory lineages, it has become self-sterile, rendering it unattractive for studies dependant on sexual reproduction. On the contrary, the more recently isolated ecotypes ‘Villersexel’ (France, 2003) and ‘Reute’ (Germany, 2006) have 15 times more sporophytes (77% of total gametophores), indicating a high fertility rate [26]. Despite these differences, all ecotypes can be propagated asexually in identical conditions from any tissue thanks to the high regeneration rate of this moss, through which explants will redifferentiate into chloronema and initiate the life cycle from there [28, 29].
Several essential developmental processes are shared between all land plants, including angiosperms and bryophytes. Strikingly, in mosses many shared processes take place with a simplified set of genes and sometimes in single cells. Therefore, the underlying genetic regulatory networks of development are easier to study. In this section, we highlight some of these processes and their unique ease of study in the moss
The ability to structure organs in three dimensions (3D) was an essential feature for water-to-land transition. Many aquatic plants develop in a homogeneous environment, whereas land plants faced a highly distinct environment at ground surface level (a plane) and at the air/soil axis (perpendicular to this plane). This required land plants to develop specific tissues to efficiently grow in each dimension and cope with new challenges [20].
From a physiological perspective, the transition from 2D-to-3D growth in mosses consists in the development of complex structures such as the gametophore that grows out of the surface plane where protonema thrive, exhibiting negative gravitropism and positive phototropism [30, 31, 32]. The development of the gametophore relies on the ability of the moss to define different organisation in each dimension of space. The basal units from which this spatial assembly takes place are ultimately single cell primordia, from which organs emanate [33]. To this end, a cell must spatially sense intrinsic and extrinsic signals and transduce them to subcellular structures that pave the way to division plane positioning and subsequent asymmetric and oriented cell division. However, the molecular basis of spatial sensing and its transduction to organisation, action and maintenance of the division machinery has not been fully elucidated yet [12].
Cellularly, the 2D-to-3D transition in a moss occurs during bud formation (Figure 1). When a cell initial is formed in protonemata, in 5% of the cases it has a bud initial cell identity instead of a branch initial identity. Each bud initial swells and undergoes divisions oriented in all three axes of space [34]. This transition is supported and maintained by a different genetic and molecular machinery than that of protonemal development, and is the molecular basis of 3D growth. These proteins are specifically present from the first cell of the bud and onwards (e.g. DEK1, NOG1), and remains active in subsequent proliferating organs (e.g. shoot apical meristem, phyllids, gametangia) [13, 34, 35, 36]. In general, gradients or local clusters of proteins, peptides, nucleic acids, and hormones can be signals, sensors and/or actuators in developmental processes, and what is upstream of the cascade of 2D-to-3D transition remains a mystery. Some actors that are already on the radar include oscillations of auxin and cytokinin concentrations, ROP and SOSEKI proteins and CLE/CLV peptides-receptors, and they have been pinned down to different moments of the process and at different locations [12, 37, 38]. However, how do they coordinate their activity remains elusive and an active area of study.
In vascular plants, the 2D-to-3D transition occurs only once in their lifecycle, during embryogenesis, where orthologs of the essential moss 3D machinery genes are also expressed [34, 35]. This event is confined in the endosperm during embryogenesis and its observation in seed plants requires seed and ovule microdissection, which makes
These features provide a privileged seat for
The protagonist of 2D-to-3D growth transition is the formation of the shoot apical meristem (SAM), a region at the apical growth side responsible for the continuous formation of the aerial organs of the plant, including leaves and reproductive organs [40, 41]. The histology of the SAM in angiosperms describes a central zone with stem cells that self-renew, divide and radially differentiate into peripheral cells that determine organ initiation at specific locations (e.g. by placement and establishment of new leaf primordia) [42]. While angiosperms present multicellular stem cell centre SAMs during their sporophytic phase, bryophytes present an unicellular structure, the shoot apical cell (SAC), both in their gametophytic and sporophytic phase (Figure 1) [43]. Despite the differences, both SAM/SAC share a common organisation, with stem cell(s) at the centre, surrounded by regularly differentiating tissue [44].
The mechanisms that establish and maintain these pluripotent stem cell(s) in the SAM/SAC is unknown. It involves spatial sensing, cell-to-cell communication, and asymmetric and oriented cell divisions, which render mosses attractive systems for their accessibility.
The developmental origin of the sporophytic SAC in
The genetic make-up in both taxa has shown to rely on auxin response through AUXIN RESPONSE FACTORS (ARFs), cytokinin signalling by ARABIDOPSIS RESPONSE REGULATORS (ARRs), CHASE domain-containing histidine kinases (CHKs) and CYTOKININ OXIDASE/DEHYDROGENASE (CKX), local coordination through several transcription factors families like CLAVATAs (CLVs), CUP-SHAPED COTELYDONS (CUC), LATERAL ORGAN BOUNDARY (LOB) and signalling peptides like CLAVATA3/EMBRYO SURROUNDING REGIONRELATED (CLE) and chromatin modification by Polycomb Repressive Complex 1 and 2 (PRC1,2), among others [48]. Although many key factors have conserved roles in SAM formation and maintenance in seed plants and mosses, some important factors in angiosperms, like the key regulator of stem cell maintenance WUSCHEL, are not found in
The most noticeable outcome of the shoot apical meristem/cell (SAM/SAC) activity is the organised and oriented initiation of leaf primordia along the stem, which leads to a unique geometric pattern of leaves and shoot branches named
However, the pattern arises from essentially different SAMs/SACs: in angiosperms, phyllotaxis derives from a multicellular system with well-reported oscillating auxin peaks around the SAM growth axis, whereas
The limited understanding of the origin and underlying molecular mechanisms of this rotating pattern and derived phyllotactic pattern is largely confined to the sporophyte of the evolutionarily recent group of flowering plants (angiosperms). The available transcriptomic data of bud and tip cells and gametophores (or 3D shoots) may provide more insight in the transition from uniplanar to triplanar meristematic growth in moss [48].
Aligned with the phenotypic similarities of moss and angiosperm phyllotactic patterns, several factors known from Arabidopsis have also been found in mosses, including receptor signalling genes involved in shoot meristem size and patterning, hormone biosynthesis genes, transcription factors that control cell-specific mechanism of developmental pattering, chromatin remodelling complexes and cell cycle [48]. Many of these factors are essentially executive and likely controlled by some spatial sensing machinery. Comparing them with new contributors or absent members in the minimal regulatory network of mosses may help unravelling the fundamental elements that trigger the orientation-specification machinery that greatly impacts plant architecture in all land plants, including relevant crops.
Most described processes in this chapter require cell identity acquisition and maintenance. In certain circumstances (e.g. wounding), differentiated plant cells can reprogram to become new stem cells, divide and redifferentiate for organ
In most tissue cultures of other plant species, exogenous hormones (e.g. auxin and cytokinin) are required to induce callus formation and plant regeneration [60]. However, in moss, cells are capable of regenerating from protoplasts or excised phyllids into new protonema filaments in the absence of exogenous hormones (Figure 1) [61]. This implies that the whole regeneration toolkit is present in mosses and can be endogenously activated on demand, which makes them different from other taxa (e.g. angiosperms) [62].
When a phyllid is excised, cells neighbouring the cutting edge can reprogram from somatic cells to protonemal stem cells, which can then start a new life cycle [14]. This regeneration process is easy to study in mosses for several reasons: firstly, cell identity conversion can be easily tracked with protonema stem cell reporters [29]; secondly, aside of the simplicity of
The mechanistic studies of the cell fate acquisition can benefit from this simple cell type conversion in comparison to other model systems used in cell reprogramming investigation (e.g. regenerative callus or Arabidopsis roots that consist of multiple cell types that possess different tissue identity) for its minimality and event frequency [65, 66]. In angiosperms, regeneration is often reduced to localised stem cell pools (e.g. the base of leaves), takes longer to establish, and it is multicellular and asynchronous at the explant level [58, 67].
Previous studies have taken advantage of the abovementioned features to investigate gene expression profile during phyllid cell reprogramming, which revealed that genes involved in stress, proteolysis, and hormone signalling pathways are induced from 6 to 24 h after cutting [64]. Some genes have been demonstrated to play essential roles in moss leaf reprogramming, including
One interesting feature of moss regeneration is inhibition of neighbouring cells. The necessary cell–cell (apoplastic e.g. Ca2+-mediated) or cell-to-cell (plasmodesmata-mediated) communication makes regeneration an attractive developmental process to study this cell crosstalk [71, 72, 73]. The phytohormone ABA is a key responsible of the dynamic regulation of the permeability of plasmodesmata in response to changing environments, such as wounding. Control in plasmodesmata pore size can influence the signalling molecules that can pass through or can be blocked in particular cells, which can have a direct effect in development of the processes mentioned until now [74, 75].
The signalling pathways and functions of plant hormones are substantially conserved in
Three plant hormones—auxin, cytokinin and strigolactone—have shown to regulate shoot branching patterns (phyllotaxis) and activation in angiosperms. Auxin moves down the main shoot of angiosperms to inhibit branch development, while cytokinin promotes branching. In addition to branching, auxin is the key molecule in the control of plant growth and development, and promotes organ differentiation [76, 77]. Exogenous application of auxin or its inhibitors results in irregular cell shapes and inhibit lateral organ formation, for instance in shoot apical meristem (SAM) maintenance. The understanding of hormone regulation and signalling in angiosperms progresses slowly due to tissue and gene network complexity. In
Cytokinins and strigolactones also influence plant architecture, both in angiosperms and mosses. The levels of cytokinin are high and precisely distributed in the central stem cell region of SAM in angiosperms to maintain stemcellness [84]. In the root apical meristem, auxin and cytokinin act antagonistically in meristem size control, but its levels, distribution and interaction in
As shown, many processes that define plant architecture are regulated in similar ways both in angiosperms and mosses. However, mosses offer a reduced gene network and regulation complexity that facilitates the analysis of hormone functions in the related developmental processes. Furthermore, subcellular and tissue-level transport and distribution of hormones can be best visualised in their simple plant bodies. In such plant models, new hormone functions will prove to be easier to study and translate to agronomically relevant plants.
Many valuable online resources with information on protocols, stocks, tools and genetic information have been exhaustively compiled elsewhere [11]. Hereby, we provide some additional information and summary of the essentials of research in
The small size and simple architecture of moss organs allows detailed microscopic visualisation easy to accomplish in almost all tissues. The strings of cells in protonema and their branching is trivial to closely visualise, and the transition to gametophores can be well tracked until the stem-like centre becomes slightly thicker than a dozen of cells and grows out of the plane. From it, the leaf-like structures (phyllids) have only one cell of thickness except in the midrib and can be tore apart for up-close visualisation. The terminal sexual organs (antheridia and archegonia) have a 3D structure that is easy to fully dissect or directly visualise due to their monolayered sack structures. The subsequently developed spore-bearing containers (sporangia) are full of single-cell spores [4].
This miniaturised body renders mosses accessible systems for the study and dissection of cell and molecular biology that require close monitoring in time and space. Such studies greatly benefit of accessibility to single cells in their context for observation of subcellular responses
In
Visualised | Fusion/Target | Purpose | Reference |
---|---|---|---|
Nucleus | NLS4-GFP-GUS | Nuclear localisation | [21] |
Endomembrane system | Endoplasmic reticulum | [11] | |
α-1,2-mannosidase | Golgi apparatus imaging | [88] | |
Targeting signal type 1 (SKL) | Peroxisome imaging | [88] | |
Mitochondria | Cytochrome c oxidase | Mitochondria imaging | [88] |
Cytoskeleton | LifeAct | Actin cytoskeleton | [89] |
Tubulin α | Microtubule cytoskeleton | [90] | |
MAP65 | Antiparallel Microtubule-microtubule contacts | [91] | |
Kinesins | [90] | ||
Plasma membrane | SNAP-TM-mCherry | Membrane tracking | [21] |
Auxin | pDR5v2:GFP-GUS | Aux. induced fluor. | [92] |
GH3:GFP-GUS | Aux. induced fluor. | ||
R2D2 | Ratiometric induction | ||
Protonema | pRM09:NLS4-GFP-GUS | Protonema identity reporting | [29] |
pRM55:NLS4-GFP-GUS | Protonema identity reporting |
Compilation of key molecular reporter lines published in literature to study cell and developmental processes in
Despite the advantageous physiological features of
Microfluidic devices are transparent and flexible structures commonly produced using the biocompatible and air-permeable polydimethylsiloxane (PDMS) polymer. In biology, they have been used for study and imaging of cell and tissue development
Until now,
Some groups carried out forward genetic screening by X-ray or chemical mutagens that generated mutants with hormone resistance or abnormal tropic responses [85, 98]. However, due to the lack of genomic information, the disrupted genes that caused the phenotypes were never identified.
Recently, the completion of genome sequencing and the establishment of its genetic mapping tools removed the obstacles in the forward genetic screening of
With this genetic mapping resource, in the past few years, researchers started to perform forward genetic screening by treating protoplast with UV light. In such screenings, mutants with phenotypic defects were successfully obtained and the causal lesions were identified by outcrossing and whole genome sequencing. Notably, essential genes that are crucial for growth may not be identified due to the lethality of their knock-outs; therefore, conditional screening was performed to overcome this problem. After the UV light treatment, plants were cultured under different temperatures and in such conditions, plants showed growth defect only in high temperature were selected as a temperature-sensitive mutant [100].
Another screening aimed to discover genes that are essential for the 2D-to-3D transition is also limited by developmental defects, given that mutants in this process cannot produce gametophores necessary for sexual crossing. To overcome this problem, instead of crossing, researchers generated somatic hybrids between Villersexel mutants and Gransden wildtype, which produced diploid sporophytes [34]. Spores released from this hybrid sporophyte exhibited consistent phenotypic segregation ratio with meiosis. Mutant plants generated from these diploid spores were sequenced and genomically mapped to achieve the identification of new crucial genes for moss 2D-to-3D transition (e.g. NO GAMETOPHORES 1 and 2, or NOG1, NOG2).
In addition to UV light, tobacco Tnt1 retrotransposon was used to produce insertional mutations in genic and GC-rich regions [101]. Both PEG- or Agrobacterium-mediated transformations were applied and successfully produced mutants.
The integration of DNA constructs (including promoter, gene of interest and selection cassettes) necessary to produce transformants with stable expression and non-disrupting phenotypes requires targeting of neutral loci that do not intrinsically produce a phenotype when disrupted, often due to gene redundancy. In Table 2, there are several standard neutral loci indicated which reportedly showed no visible phenotypes or morphological defects when it is replaced by an entire gene expression cassette.
Locus | Vector | Purpose | Reference |
---|---|---|---|
PIG1 locus | pGX8 pGG626 | XVE inducible overexpression XVE inducible RNAi expression | [102, 103] |
Pp108 locus | pUGGi | Constitutive RNAi expression | [104] |
Pp108 locus | pTH-Ubi-Gate | Constitutive expression by the maize ubiquitin promoter | [105] |
Redundant copy of the ARPC2 gene | pTK-Ubi-Gate | ||
Redundant copy of the ARPC3 gene | pTZ-Ubi-Gate | ||
PTA1 locus | pT10G | Overexpression by EF1α promoter | [106] |
BS213 locus | pMJ1 | [107] |
A compilation of vectors designed to target proven neutral loci in
Currently there are several vector sets released to specifically target neutral loci, that contain their flanking regions homologous to parts of the locus at start and end of the vector. Cloning the gene of interest in between readily allows replacement of the targeted locus with the entire cassette via homologous recombination (see section
In
The game-changing CRISPR/Cas9 method has proven an efficient and effective tool in
In Nogué’s lab, a co-delivery method was developed where each element of the system (Cas9, sgRNA and selection cassettes) was present in a separate plasmid. In this method, Cas9 expression is driven by an actin promoter and ready to use as is. The selection strategy can be chosen freely due to the lack of integration, minding the presence of resistance in the background lines. This method is also suitable for multiple mutations in different genes at once, given that more than one sgRNA plasmid can be simultaneously delivered in one transformation with still sufficiently high efficiencies of transformation [110].
In Bezanilla’s lab, a whole set of gateway destination vectors for CRISPR/Cas9 system was developed. The strategy was to design a vector set to finally put all the three essential components (Cas9, sgRNA and selection cassettes) in a single expression plasmid. In both protocols, the sgRNA can be designed and optimised using the online design tool CRISPOR V1 against
To increase the accuracy of mutations, the CRISPR/Cas9 system is applied with a homology-directed repair (HDR), which allows for seamless knock-in or point mutation in desired sites. The template DNA can be a donor plasmid that harbours homologous fragments or oligodeoxynucleotides (ODNs) [111, 113]. By co-transforming the plasmid or ODNs together with CRISPR/Cas9 and sgRNA vectors, both methods show high accuracy to generate a desired point mutation or scarless insertion with a fluorescence tag at any suitable location of the gene.
Given that moss possesses a relatively big gene family, arguably due to its double genome duplication, the employment of gene deletion strategies might be inefficient to investigate gene functions due to the high redundancy rate (e.g. there are four ROP genes with highly homologous or identical sequences) [38]. For this, RNA interference (RNAi) strategies offer an alternative to overcome this problem, and the procedure has been well-established.
To generate an RNAi construct for a gene of interest, a DNA sequence of 300 to 1000 bp is subcloned in a destination vector. After standard PEG-mediated transformation, the silencing effect can be detected after 24 h and last up to 3 weeks [104]. To avoid lethal effects when constitutively expressing interfering RNA, an oestradiol-inducible RNAi system is available [102]. Coupling RNAi silencing activation with fluorescent reporters facilitates screenings of loss of function phenotypes [104].
Standard transformation protocols have been applicable to
Strategy | Highlights | Reference |
---|---|---|
PEG/Mannitol | One to two round selection, 10% of transformants, 3–4 weeks for stable transformants | [114, 115, 116, 117] |
Four-round selection, 100% positive transformants, 12–16 weeks | [118] | |
Particle bombardment | Easy to conduct; less used. Transient and stable DNA integration. | [119] |
Summary table of the classical and current transformation techniques and reference protocols for application.
Due to their accessibility, tractability and close yet independent phylogenetic position, the interest in Bryophytes has increased dramatically in the last decade [1]. Beyond
The fire moss
The Hypnales (Bryopsida) are the biggest and most diverse order of mosses with varied morphology, and mostly exhibit pleurocarpous (i.e. non-erect) growth fashion. It includes
Remarkably, the common aquatic moss
The genus
The moss class Polytrichopsida is the second biggest (~200 species) after Bryopsida (~11500 species), and its species have unique morphological characteristics that make them uniquely interesting for developmental biology [5]. Despite mosses being regarded as avascular plants, some exhibit hydroids and leptoids, a functionally analogous tissue to tracheids and sieve elements of vascular plants that slightly differs morphologically and developmentally. Polytrichopsida has several genera with such structures, in some cases underdeveloped and, in others, completely functional, like in
Despite its potential to provide valuable insight, genetic tools have barely been developed for this taxon, but full mitochondrial and plastid genomes have recently been published for
The desert moss
The heavy metal-tolerant moss
We have shown how mosses are an increasingly relevant model group to plant developmental biology due to their distinct accessibility at physiological and molecular level. Their evolutionary distance with agronomically relevant plants does not diminish their potential to help to understand fundamental questions of development that remain unsolved, given that most essential regulatory networks are conserved. This has been shown in the hormonal regulation of branching and shoot and root development, regeneration, the establishment of shoot apical cells and the genetic make-up in 2D-to-3D transition. The utility of mosses has convinced the scientific community to the point of promoting the sequencing of eight new species to explore other physiological processes in the last few years. We have also shown how
The authors would like to thank Dr. Jeroen de Keijzer and Dr. Tijs Ketelaar for their thoughtful and detailed review of the manuscript. Also, the funding agencies Technology, Knowledge and Innovation, division Horticulture and Propagating Material (TKI T&U) and the Dutch Research Council (NWO) (reference number: TKILWV20.390) for funding JFC and the ERC grant to Prof. J. Friml (reference number: PR1023ERC02) for funding HT. The authors would like to sincerely apologise for the literature not cited that may be relevant for this chapter and is not present due to space constraints.
"Open access contributes to scientific excellence and integrity. It opens up research results to wider analysis. It allows research results to be reused for new discoveries. And it enables the multi-disciplinary research that is needed to solve global 21st century problems. Open access connects science with society. It allows the public to engage with research. To go behind the headlines. And look at the scientific evidence. And it enables policy makers to draw on innovative solutions to societal challenges".
\n\nCarlos Moedas, the European Commissioner for Research Science and Innovation at the STM Annual Frankfurt Conference, October 2016.
",metaTitle:"About Open Access",metaDescription:"Open access contributes to scientific excellence and integrity. It opens up research results to wider analysis. It allows research results to be reused for new discoveries. And it enables the multi-disciplinary research that is needed to solve global 21st century problems. Open access connects science with society. It allows the public to engage with research. To go behind the headlines. And look at the scientific evidence. And it enables policy makers to draw on innovative solutions to societal challenges.\n\nCarlos Moedas, the European Commissioner for Research Science and Innovation at the STM Annual Frankfurt Conference, October 2016.",metaKeywords:null,canonicalURL:"about-open-access",contentRaw:'[{"type":"htmlEditorComponent","content":"The Open Access publishing movement started in the early 2000s when academic leaders from around the world participated in the formation of the Budapest Initiative. They developed recommendations for an Open Access publishing process, “which has worked for the past decade to provide the public with unrestricted, free access to scholarly research—much of which is publicly funded. Making the research publicly available to everyone—free of charge and without most copyright and licensing restrictions—will accelerate scientific research efforts and allow authors to reach a larger number of readers” (reference: http://www.budapestopenaccessinitiative.org)
\\n\\nIntechOpen’s co-founders, both scientists themselves, created the company while undertaking research in robotics at Vienna University. Their goal was to spread research freely “for scientists, by scientists’ to the rest of the world via the Open Access publishing model. The company soon became a signatory of the Budapest Initiative, which currently has more than 1000 supporting organizations worldwide, ranging from universities to funders.
\\n\\nAt IntechOpen today, we are still as committed to working with organizations and people who care about scientific discovery, to putting the academic needs of the scientific community first, and to providing an Open Access environment where scientists can maximize their contribution to scientific advancement. By opening up access to the world’s scientific research articles and book chapters, we aim to facilitate greater opportunity for collaboration, scientific discovery and progress. We subscribe wholeheartedly to the Open Access definition:
\\n\\n“By “open access” to [peer-reviewed research literature], we mean its free availability on the public internet, permitting any users to read, download, copy, distribute, print, search, or link to the full texts of these articles, crawl them for indexing, pass them as data to software, or use them for any other lawful purpose, without financial, legal, or technical barriers other than those inseparable from gaining access to the internet itself. The only constraint on reproduction and distribution, and the only role for copyright in this domain, should be to give authors control over the integrity of their work and the right to be properly acknowledged and cited” (reference: http://www.budapestopenaccessinitiative.org)
\\n\\nOAI-PMH
\\n\\nAs a firm believer in the wider dissemination of knowledge, IntechOpen supports the Open Access Initiative Protocol for Metadata Harvesting (OAI-PMH Version 2.0). Read more
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\\n\\nBook chapters published in edited volumes are distributed under the Creative Commons Attribution 3.0 Unported License (CC BY 3.0). IntechOpen upholds a very flexible Copyright Policy. There is no copyright transfer to the publisher and Authors retain exclusive copyright to their work. All Monographs/Compacts are distributed under the Creative Commons Attribution-NonCommercial 4.0 International (CC BY-NC 4.0). Read more
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\\n\\nDigital Archiving Policy
\\n\\nIntechOpen is committed to ensuring the long-term preservation and the availability of all scholarly research we publish. We employ a variety of means to enable us to deliver on our commitments to the scientific community. Apart from preservation by the Croatian National Library (for publications prior to April 18, 2018) and the British Library (for publications after April 18, 2018), our entire catalogue is preserved in the CLOCKSS archive.
\\n\\nOpen Science is transparent and accessible knowledge that is shared and developed through collaborative networks.
\\n\\nOpen Science is about increased rigour, accountability, and reproducibility for research. It is based on the principles of inclusion, fairness, equity, and sharing, and ultimately seeks to change the way research is done, who is involved and how it is valued. It aims to make research more open to participation, review/refutation, improvement and (re)use for the world to benefit.
\\n\\nOpen Science refers to doing traditional science with more transparency involved at various stages, for example by openly sharing code and data. It implies a growing set of practices - within different disciplines - aiming at:
\\n\\nWe aim at improving the quality and availability of scholarly communication by promoting and practicing:
\\n\\n\\n"}]'},components:[{type:"htmlEditorComponent",content:'
The Open Access publishing movement started in the early 2000s when academic leaders from around the world participated in the formation of the Budapest Initiative. They developed recommendations for an Open Access publishing process, “which has worked for the past decade to provide the public with unrestricted, free access to scholarly research—much of which is publicly funded. Making the research publicly available to everyone—free of charge and without most copyright and licensing restrictions—will accelerate scientific research efforts and allow authors to reach a larger number of readers” (reference: http://www.budapestopenaccessinitiative.org)
\n\nIntechOpen’s co-founders, both scientists themselves, created the company while undertaking research in robotics at Vienna University. Their goal was to spread research freely “for scientists, by scientists’ to the rest of the world via the Open Access publishing model. The company soon became a signatory of the Budapest Initiative, which currently has more than 1000 supporting organizations worldwide, ranging from universities to funders.
\n\nAt IntechOpen today, we are still as committed to working with organizations and people who care about scientific discovery, to putting the academic needs of the scientific community first, and to providing an Open Access environment where scientists can maximize their contribution to scientific advancement. By opening up access to the world’s scientific research articles and book chapters, we aim to facilitate greater opportunity for collaboration, scientific discovery and progress. We subscribe wholeheartedly to the Open Access definition:
\n\n“By “open access” to [peer-reviewed research literature], we mean its free availability on the public internet, permitting any users to read, download, copy, distribute, print, search, or link to the full texts of these articles, crawl them for indexing, pass them as data to software, or use them for any other lawful purpose, without financial, legal, or technical barriers other than those inseparable from gaining access to the internet itself. The only constraint on reproduction and distribution, and the only role for copyright in this domain, should be to give authors control over the integrity of their work and the right to be properly acknowledged and cited” (reference: http://www.budapestopenaccessinitiative.org)
\n\nOAI-PMH
\n\nAs a firm believer in the wider dissemination of knowledge, IntechOpen supports the Open Access Initiative Protocol for Metadata Harvesting (OAI-PMH Version 2.0). Read more
\n\nLicense
\n\nBook chapters published in edited volumes are distributed under the Creative Commons Attribution 3.0 Unported License (CC BY 3.0). IntechOpen upholds a very flexible Copyright Policy. There is no copyright transfer to the publisher and Authors retain exclusive copyright to their work. All Monographs/Compacts are distributed under the Creative Commons Attribution-NonCommercial 4.0 International (CC BY-NC 4.0). Read more
\n\nPeer Review Policies
\n\nAll scientific works are Peer Reviewed prior to publishing. Read more
\n\nOA Publishing Fees
\n\nThe Open Access publishing model employed by IntechOpen eliminates subscription charges and pay-per-view fees, enabling readers to access research at no cost. In order to sustain operations and keep our publications freely accessible we levy an Open Access Publishing Fee for manuscripts, which helps us cover the costs of editorial work and the production of books. Read more
\n\nDigital Archiving Policy
\n\nIntechOpen is committed to ensuring the long-term preservation and the availability of all scholarly research we publish. We employ a variety of means to enable us to deliver on our commitments to the scientific community. Apart from preservation by the Croatian National Library (for publications prior to April 18, 2018) and the British Library (for publications after April 18, 2018), our entire catalogue is preserved in the CLOCKSS archive.
\n\nOpen Science is transparent and accessible knowledge that is shared and developed through collaborative networks.
\n\nOpen Science is about increased rigour, accountability, and reproducibility for research. It is based on the principles of inclusion, fairness, equity, and sharing, and ultimately seeks to change the way research is done, who is involved and how it is valued. It aims to make research more open to participation, review/refutation, improvement and (re)use for the world to benefit.
\n\nOpen Science refers to doing traditional science with more transparency involved at various stages, for example by openly sharing code and data. It implies a growing set of practices - within different disciplines - aiming at:
\n\nWe aim at improving the quality and availability of scholarly communication by promoting and practicing:
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Among different technologies for hydrogen production, oxygenic natural and artificial photosynthesis using direct photochemistry in synthetic complexes have a great potential to produce hydrogen as both use clean and cheap sources - water and solar energy. Photosynthetic organisms capture sunlight very efficiently and convert it into organic molecules. Artificial photosynthesis is one way to produce hydrogen from water using sunlight by employing biomimetic complexes. However, splitting of water into protons and oxygen is energetically demanding and chemically difficult. In oxygenic photosynthetic microorganisms water is splitted into electrons and protons during primary photosynthetic processes. The electrons and protons are redirected through the photosynthetic electron transport chain to the hydrogen-producing enzymes-hydrogenase or nitrogenase. By these enzymes, e- and H+ recombine and form gaseous hydrogen. Biohydrogen activity of hydrogenase can be very high but it is extremely sensitive to photosynthetic O2. At the moment, the efficiency of biohydrogen production is low. However, theoretical expectations suggest that the rates of photon conversion efficiency for H2 bioproduction can be high enough (> 10%). Our review examines the main pathways of H2 photoproduction using photosynthetic organisms and biomimetic photosynthetic systems and focuses on developing new technologies based on the effective principles of photosynthesis.",book:{id:"3587",slug:"biomimetics-learning-from-nature",title:"Biomimetics",fullTitle:"Biomimetics Learning from Nature"},signatures:"Suleyman I. Allakhverdiev, Vladimir D. Kreslavski, Velmurugan Thavasi, Sergei K. Zharmukhamedov, Vyacheslav V. 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It performs very complex tasks while occupying about 2 liters of volume and consuming very little energy. The computation tasks are performed by special cells in the brain called neurons. They compute using electrical pulses and exchange information between them through chemicals called neurotransmitters. With this as inspiration, there are several compute models which exist today trying to exploit the inherent efficiencies demonstrated by nature. The compute models representing spiking neural networks (SNNs) are biologically plausible, hence are used to study and understand the workings of brain and nervous system. More importantly, they are used to solve a wide variety of problems in the field of artificial intelligence (AI). They are uniquely suited to model temporal and spatio-temporal data paradigms. 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Finally, applications on the chaotic time series prediction and the video frame recognition will be demonstrated.",book:{id:"6875",slug:"bio-inspired-technology",title:"Bio-Inspired Technology",fullTitle:"Bio-Inspired Technology"},signatures:"Kangjun Bai and Yang Yi",authors:[{id:"239041",title:"Prof.",name:"Yang",middleName:null,surname:"Yi",slug:"yang-yi",fullName:"Yang Yi"},{id:"245542",title:"Mr.",name:"Kangjun",middleName:null,surname:"Bai",slug:"kangjun-bai",fullName:"Kangjun Bai"}]},{id:"58622",title:"Bio-inspired Adaptable Facade Control Reflecting User's Behavior",slug:"bio-inspired-adaptable-facade-control-reflecting-user-s-behavior",totalDownloads:1619,totalCrossrefCites:0,totalDimensionsCites:1,abstract:"The purpose of this research is to develop the process of methodology in designing adaptable façade. This study focuses on the processes of façade operation control for each resident’s unit according to the user’s lifestyle. 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His research focuses on biochemistry, biophysics, genetics, molecular biology, and molecular medicine with specialization in the fields of drug design, protein structure-function, protein folding, prions, microRNA, pseudogenes, molecular cancer, epigenetics, metabolites, proteomics, genomics, protein expression, and characterization by spectroscopic and calorimetric methods.",institutionString:"University of Health Sciences",institution:null},{id:"180528",title:"Dr.",name:"Hiroyuki",middleName:null,surname:"Kagechika",slug:"hiroyuki-kagechika",fullName:"Hiroyuki Kagechika",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/180528/images/system/180528.jpg",biography:"Hiroyuki Kagechika received his bachelor’s degree and Ph.D. in Pharmaceutical Sciences from the University of Tokyo, Japan, where he served as an associate professor until 2004. He is currently a professor at the Institute of Biomaterials and Bioengineering (IBB), Tokyo Medical and Dental University (TMDU). From 2010 to 2012, he was the dean of the Graduate School of Biomedical Science. Since 2012, he has served as the vice dean of the Graduate School of Medical and Dental Sciences. He has been the director of the IBB since 2020. Dr. Kagechika’s major research interests are the medicinal chemistry of retinoids, vitamins D/K, and nuclear receptors. 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Then, he was appointed as an Assistant Professor of Neurology, UTHSC in 2005. He moved to the Cleveland Clinic in USA as a Project Scientist/Staff in 2006 where he focused on the studies of eye disease proteomics and biomarkers. He returned to UTHSC as an Assistant Professor of Neurology in the end of 2007, engaging in proteomics and biomarker studies of lung diseases and brain tumors, and initiating the studies of predictive, preventive, and personalized medicine (PPPM) in cancer. In 2010, he was promoted to Associate Professor of Neurology, UTHSC. Currently, he is a Professor at Xiangya Hospital of Central South University in China, Fellow of Royal Society of Medicine (FRSM), the European EPMA National Representative in China, Regular Member of American Association for the Advancement of Science (AAAS), European Cooperation of Science and Technology (e-COST) grant evaluator, Associate Editors of BMC Genomics, BMC Medical Genomics, EPMA Journal, and Frontiers in Endocrinology, Executive Editor-in-Chief of Med One. He has\npublished 116 peer-reviewed research articles, 16 book chapters, 2 books, and 2 US patents. 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He has published several articles in peer-reviewed journals, chapters, and edited books. His area of specialization is free radical biochemistry and autoimmune diseases.",institutionString:"Imam Abdulrahman Bin Faisal University",institution:{name:"Imam Abdulrahman Bin Faisal University",country:{name:"Saudi Arabia"}}},{id:"41865",title:"Prof.",name:"Farid A.",middleName:null,surname:"Badria",slug:"farid-a.-badria",fullName:"Farid A. Badria",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/41865/images/system/41865.jpg",biography:"Farid A. Badria, Ph.D., is the recipient of several awards, including The World Academy of Sciences (TWAS) Prize for Public Understanding of Science; the World Intellectual Property Organization (WIPO) Gold Medal for best invention; Outstanding Arab Scholar, Kuwait; and the Khwarizmi International Award, Iran. He has 250 publications, 12 books, 20 patents, and several marketed pharmaceutical products to his credit. He continues to lead research projects on developing new therapies for liver, skin disorders, and cancer. Dr. Badria was listed among the world’s top 2% of scientists in medicinal and biomolecular chemistry in 2019 and 2020. He is a member of the Arab Development Fund, Kuwait; International Cell Research Organization–United Nations Educational, Scientific and Cultural Organization (ICRO–UNESCO), Chile; and UNESCO Biotechnology France",institutionString:"Mansoura University",institution:{name:"Mansoura University",country:{name:"Egypt"}}},{id:"329385",title:"Dr.",name:"Rajesh K.",middleName:"Kumar",surname:"Singh",slug:"rajesh-k.-singh",fullName:"Rajesh K. Singh",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/329385/images/system/329385.png",biography:"Dr. Singh received a BPharm (2003) and MPharm (2005) from Panjab University, Chandigarh, India, and a Ph.D. (2013) from Punjab Technical University (PTU), Jalandhar, India. He has more than sixteen years of teaching experience and has supervised numerous postgraduate and Ph.D. students. He has to his credit more than seventy papers in SCI- and SCOPUS-indexed journals, fifty-five conference proceedings, four books, six Best Paper Awards, and five projects from different government agencies. He is currently an editorial board member of eight international journals and a reviewer for more than fifty scientific journals. He received Top Reviewer and Excellent Peer Reviewer Awards from Publons in 2016 and 2017, respectively. He is also on the panel of The International Reviewer for reviewing research proposals for grants from the Royal Society. He also serves as a Publons Academy mentor and Bentham brand ambassador.",institutionString:"Punjab Technical University",institution:{name:"Punjab Technical University",country:{name:"India"}}},{id:"142388",title:"Dr.",name:"Thiago",middleName:"Gomes",surname:"Gomes Heck",slug:"thiago-gomes-heck",fullName:"Thiago Gomes Heck",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/142388/images/7259_n.jpg",biography:null,institutionString:null,institution:{name:"Universidade Regional do Noroeste do Estado do Rio Grande do Sul",country:{name:"Brazil"}}},{id:"336273",title:"Assistant Prof.",name:"Janja",middleName:null,surname:"Zupan",slug:"janja-zupan",fullName:"Janja Zupan",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/336273/images/14853_n.jpeg",biography:"Janja Zupan graduated in 2005 at the Department of Clinical Biochemistry (superviser prof. dr. Janja Marc) in the field of genetics of osteoporosis. Since November 2009 she is working as a Teaching Assistant at the Faculty of Pharmacy, Department of Clinical Biochemistry. In 2011 she completed part of her research and PhD work at Institute of Genetics and Molecular Medicine, University of Edinburgh. She finished her PhD entitled The influence of the proinflammatory cytokines on the RANK/RANKL/OPG in bone tissue of osteoporotic and osteoarthritic patients in 2012. From 2014-2016 she worked at the Institute of Biomedical Sciences, University of Aberdeen as a postdoctoral research fellow on UK Arthritis research project where she gained knowledge in mesenchymal stem cells and regenerative medicine. She returned back to University of Ljubljana, Faculty of Pharmacy in 2016. She is currently leading project entitled Mesenchymal stem cells-the keepers of tissue endogenous regenerative capacity facing up to aging of the musculoskeletal system funded by Slovenian Research Agency.",institutionString:null,institution:{name:"University of Ljubljana",country:{name:"Slovenia"}}},{id:"357453",title:"Dr.",name:"Radheshyam",middleName:null,surname:"Maurya",slug:"radheshyam-maurya",fullName:"Radheshyam Maurya",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/357453/images/16535_n.jpg",biography:null,institutionString:null,institution:{name:"University of Hyderabad",country:{name:"India"}}},{id:"418340",title:"Dr.",name:"Jyotirmoi",middleName:null,surname:"Aich",slug:"jyotirmoi-aich",fullName:"Jyotirmoi Aich",position:null,profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0033Y000038Ugi5QAC/Profile_Picture_2022-04-15T07:48:28.png",biography:"Biotechnologist with 15 years of research including 6 years of teaching experience. Demonstrated record of scientific achievements through consistent publication record (H index = 13, with 874 citations) in high impact journals such as Nature Communications, Oncotarget, Annals of Oncology, PNAS, and AJRCCM, etc. Strong research professional with a post-doctorate from ACTREC where I gained experimental oncology experience in clinical settings and a doctorate from IGIB where I gained expertise in asthma pathophysiology. A well-trained biotechnologist with diverse experience on the bench across different research themes ranging from asthma to cancer and other infectious diseases. An individual with a strong commitment and innovative mindset. Have the ability to work on diverse projects such as regenerative and molecular medicine with an overall mindset of improving healthcare.",institutionString:"DY Patil Deemed to Be University",institution:null},{id:"349288",title:"Prof.",name:"Soumya",middleName:null,surname:"Basu",slug:"soumya-basu",fullName:"Soumya Basu",position:null,profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0033Y000035QxIDQA0/Profile_Picture_2022-04-15T07:47:01.jpg",biography:"Soumya Basu, Ph.D., is currently working as an Associate Professor at Dr. D. Y. Patil Biotechnology and Bioinformatics Institute, Dr. D. Y. Patil Vidyapeeth, Pune, Maharashtra, India. With 16+ years of trans-disciplinary research experience in Drug Design, development, and pre-clinical validation; 20+ research article publications in journals of repute, 9+ years of teaching experience, trained with cross-disciplinary education, Dr. Basu is a life-long learner and always thrives for new challenges.\r\nHer research area is the design and synthesis of small molecule partial agonists of PPAR-γ in lung cancer. She is also using artificial intelligence and deep learning methods to understand the exosomal miRNA’s role in cancer metastasis. Dr. Basu is the recipient of many awards including the Early Career Research Award from the Department of Science and Technology, Govt. of India. She is a reviewer of many journals like Molecular Biology Reports, Frontiers in Oncology, RSC Advances, PLOS ONE, Journal of Biomolecular Structure & Dynamics, Journal of Molecular Graphics and Modelling, etc. She has edited and authored/co-authored 21 journal papers, 3 book chapters, and 15 abstracts. She is a Board of Studies member at her university. She is a life member of 'The Cytometry Society”-in India and 'All India Cell Biology Society”- in India.",institutionString:"Dr. D.Y. Patil Vidyapeeth, Pune",institution:{name:"Dr. D.Y. Patil Vidyapeeth, Pune",country:{name:"India"}}},{id:"354817",title:"Dr.",name:"Anubhab",middleName:null,surname:"Mukherjee",slug:"anubhab-mukherjee",fullName:"Anubhab Mukherjee",position:null,profilePictureURL:"https://intech-files.s3.amazonaws.com/0033Y0000365PbRQAU/ProfilePicture%202022-04-15%2005%3A11%3A18.480",biography:"A former member of Laboratory of Nanomedicine, Brigham and Women’s Hospital, Harvard University, Boston, USA, Dr. Anubhab Mukherjee is an ardent votary of science who strives to make an impact in the lives of those afflicted with cancer and other chronic/acute ailments. He completed his Ph.D. from CSIR-Indian Institute of Chemical Technology, Hyderabad, India, having been skilled with RNAi, liposomal drug delivery, preclinical cell and animal studies. He pursued post-doctoral research at College of Pharmacy, Health Science Center, Texas A & M University and was involved in another postdoctoral research at Department of Translational Neurosciences and Neurotherapeutics, John Wayne Cancer Institute, Santa Monica, California. In 2015, he worked in Harvard-MIT Health Sciences & Technology as a visiting scientist. He has substantial experience in nanotechnology-based formulation development and successfully served various Indian organizations to develop pharmaceuticals and nutraceutical products. He is an inventor in many US patents and an author in many peer-reviewed articles, book chapters and books published in various media of international repute. Dr. Mukherjee is currently serving as Principal Scientist, R&D at Esperer Onco Nutrition (EON) Pvt. Ltd. and heads the Hyderabad R&D center of the organization.",institutionString:"Esperer Onco Nutrition Pvt Ltd.",institution:null},{id:"319365",title:"Assistant Prof.",name:"Manash K.",middleName:null,surname:"Paul",slug:"manash-k.-paul",fullName:"Manash K. Paul",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/319365/images/system/319365.png",biography:"Manash K. Paul is a Principal Investigator and Scientist at the University of California Los Angeles. He has contributed significantly to the fields of stem cell biology, regenerative medicine, and lung cancer. His research focuses on various signaling processes involved in maintaining stem cell homeostasis during the injury-repair process, deciphering lung stem cell niche, pulmonary disease modeling, immuno-oncology, and drug discovery. He is currently investigating the role of extracellular vesicles in premalignant lung cell migration and detecting the metastatic phenotype of lung cancer via machine-learning-based analyses of exosomal signatures. Dr. Paul has published in more than fifty peer-reviewed international journals and is highly cited. He is the recipient of many awards, including the UCLA Vice Chancellor’s award, a senior member of the Institute of Electrical and Electronics Engineers (IEEE), and an editorial board member for several international journals.",institutionString:"University of California Los Angeles",institution:{name:"University of California Los Angeles",country:{name:"United States of America"}}},{id:"311457",title:"Dr.",name:"Júlia",middleName:null,surname:"Scherer Santos",slug:"julia-scherer-santos",fullName:"Júlia Scherer Santos",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/311457/images/system/311457.jpg",biography:"Dr. Júlia Scherer Santos works in the areas of cosmetology, nanotechnology, pharmaceutical technology, beauty, and aesthetics. Dr. Santos also has experience as a professor of graduate courses. Graduated in Pharmacy, specialization in Cosmetology and Cosmeceuticals applied to aesthetics, specialization in Aesthetic and Cosmetic Health, and a doctorate in Pharmaceutical Nanotechnology. Teaching experience in Pharmacy and Aesthetics and Cosmetics courses. She works mainly on the following subjects: nanotechnology, cosmetology, pharmaceutical technology, aesthetics.",institutionString:"Universidade Federal de Juiz de Fora",institution:{name:"Universidade Federal de Juiz de Fora",country:{name:"Brazil"}}},{id:"219081",title:"Dr.",name:"Abdulsamed",middleName:null,surname:"Kükürt",slug:"abdulsamed-kukurt",fullName:"Abdulsamed Kükürt",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/219081/images/system/219081.png",biography:"Dr. Kükürt graduated from Uludağ University in Turkey. He started his academic career as a Research Assistant in the Department of Biochemistry at Kafkas University. In 2019, he completed his Ph.D. program in the Department of Biochemistry at the Institute of Health Sciences. He is currently working at the Department of Biochemistry, Kafkas University. He has 27 published research articles in academic journals, 11 book chapters, and 37 papers. He took part in 10 academic projects. He served as a reviewer for many articles. He still serves as a member of the review board in many academic journals.",institutionString:"Kafkas University",institution:{name:"Kafkas University",country:{name:"Turkey"}}},{id:"178366",title:"Associate Prof.",name:"Volkan",middleName:null,surname:"Gelen",slug:"volkan-gelen",fullName:"Volkan Gelen",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/178366/images/system/178366.jpg",biography:"Volkan Gelen is a Physiology specialist who received his veterinary degree from Kafkas University in 2011. Between 2011-2015, he worked as an assistant at Atatürk University, Faculty of Veterinary Medicine, Department of Physiology. In 2016, he joined Kafkas University, Faculty of Veterinary Medicine, Department of Physiology as an assistant professor. Dr. Gelen has been engaged in various academic activities at Kafkas University since 2016. There he completed 5 projects and has 3 ongoing projects. He has 60 articles published in scientific journals and 20 poster presentations in scientific congresses. His research interests include physiology, endocrine system, cancer, diabetes, cardiovascular system diseases, and isolated organ bath system studies.",institutionString:"Kafkas University",institution:{name:"Kafkas University",country:{name:"Turkey"}}},{id:"418963",title:"Dr.",name:"Augustine Ododo",middleName:"Augustine",surname:"Osagie",slug:"augustine-ododo-osagie",fullName:"Augustine Ododo Osagie",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/418963/images/16900_n.jpg",biography:"Born into the family of Osagie, a prince of the Benin Kingdom. I am currently an academic in the Department of Medical Biochemistry, University of Benin. Part of the duties are to teach undergraduate students and conduct academic research.",institutionString:null,institution:{name:"University of Benin",country:{name:"Nigeria"}}},{id:"192992",title:"Prof.",name:"Shagufta",middleName:null,surname:"Perveen",slug:"shagufta-perveen",fullName:"Shagufta Perveen",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/192992/images/system/192992.png",biography:"Prof. Shagufta Perveen is a Distinguish Professor in the Department of Pharmacognosy, College of Pharmacy, King Saud University, Riyadh, Saudi Arabia. Dr. Perveen has acted as the principal investigator of major research projects funded by the research unit of King Saud University. She has more than ninety original research papers in peer-reviewed journals of international repute to her credit. She is a fellow member of the Royal Society of Chemistry UK and the American Chemical Society of the United States.",institutionString:"King Saud University",institution:{name:"King Saud University",country:{name:"Saudi Arabia"}}},{id:"49848",title:"Dr.",name:"Wen-Long",middleName:null,surname:"Hu",slug:"wen-long-hu",fullName:"Wen-Long Hu",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/49848/images/system/49848.jpg",biography:"Wen-Long Hu is Chief of the Division of Acupuncture, Department of Chinese Medicine at Kaohsiung Chang Gung Memorial Hospital, as well as an adjunct associate professor at Fooyin University and Kaohsiung Medical University. Wen-Long is President of Taiwan Traditional Chinese Medicine Medical Association. He has 28 years of experience in clinical practice in laser acupuncture therapy and 34 years in acupuncture. He is an invited speaker for lectures and workshops in laser acupuncture at many symposiums held by medical associations. He owns the patent for herbal preparation and producing, and for the supercritical fluid-treated needle. Dr. Hu has published three books, 12 book chapters, and more than 30 papers in reputed journals, besides serving as an editorial board member of repute.",institutionString:"Kaohsiung Chang Gung Memorial Hospital",institution:{name:"Kaohsiung Chang Gung Memorial Hospital",country:{name:"Taiwan"}}},{id:"298472",title:"Prof.",name:"Andrey V.",middleName:null,surname:"Grechko",slug:"andrey-v.-grechko",fullName:"Andrey V. Grechko",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/298472/images/system/298472.png",biography:"Andrey Vyacheslavovich Grechko, Ph.D., Professor, is a Corresponding Member of the Russian Academy of Sciences. He graduated from the Semashko Moscow Medical Institute (Semashko National Research Institute of Public Health) with a degree in Medicine (1998), the Clinical Department of Dermatovenerology (2000), and received a second higher education in Psychology (2009). Professor A.V. Grechko held the position of Сhief Physician of the Central Clinical Hospital in Moscow. He worked as a professor at the faculty and was engaged in scientific research at the Medical University. Starting in 2013, he has been the initiator of the creation of the Federal Scientific and Clinical Center for Intensive Care and Rehabilitology, Moscow, Russian Federation, where he also serves as Director since 2015. He has many years of experience in research and teaching in various fields of medicine, is an author/co-author of more than 200 scientific publications, 13 patents, 15 medical books/chapters, including Chapter in Book «Metabolomics», IntechOpen, 2020 «Metabolomic Discovery of Microbiota Dysfunction as the Cause of Pathology».",institutionString:"Federal Research and Clinical Center of Intensive Care Medicine and Rehabilitology",institution:null},{id:"199461",title:"Prof.",name:"Natalia V.",middleName:null,surname:"Beloborodova",slug:"natalia-v.-beloborodova",fullName:"Natalia V. Beloborodova",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/199461/images/system/199461.jpg",biography:'Natalia Vladimirovna Beloborodova was educated at the Pirogov Russian National Research Medical University, with a degree in pediatrics in 1980, a Ph.D. in 1987, and a specialization in Clinical Microbiology from First Moscow State Medical University in 2004. She has been a Professor since 1996. Currently, she is the Head of the Laboratory of Metabolism, a division of the Federal Research and Clinical Center of Intensive Care Medicine and Rehabilitology, Moscow, Russian Federation. N.V. Beloborodova has many years of clinical experience in the field of intensive care and surgery. She studies infectious complications and sepsis. She initiated a series of interdisciplinary clinical and experimental studies based on the concept of integrating human metabolism and its microbiota. Her scientific achievements are widely known: she is the recipient of the Marie E. Coates Award \\"Best lecturer-scientist\\" Gustafsson Fund, Karolinska Institutes, Stockholm, Sweden, and the International Sepsis Forum Award, Pasteur Institute, Paris, France (2014), etc. Professor N.V. Beloborodova wrote 210 papers, five books, 10 chapters and has edited four books.',institutionString:"Federal Research and Clinical Center of Intensive Care Medicine and Rehabilitology",institution:null},{id:"354260",title:"Ph.D.",name:"Tércio Elyan",middleName:"Azevedo",surname:"Azevedo Martins",slug:"tercio-elyan-azevedo-martins",fullName:"Tércio Elyan Azevedo Martins",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/354260/images/16241_n.jpg",biography:"Graduated in Pharmacy from the Federal University of Ceará with the modality in Industrial Pharmacy, Specialist in Production and Control of Medicines from the University of São Paulo (USP), Master in Pharmaceuticals and Medicines from the University of São Paulo (USP) and Doctor of Science in the program of Pharmaceuticals and Medicines by the University of São Paulo. Professor at Universidade Paulista (UNIP) in the areas of chemistry, cosmetology and trichology. Assistant Coordinator of the Higher Course in Aesthetic and Cosmetic Technology at Universidade Paulista Campus Chácara Santo Antônio. Experience in the Pharmacy area, with emphasis on Pharmacotechnics, Pharmaceutical Technology, Research and Development of Cosmetics, acting mainly on topics such as cosmetology, antioxidant activity, aesthetics, photoprotection, cyclodextrin and thermal analysis.",institutionString:null,institution:{name:"University of Sao Paulo",country:{name:"Brazil"}}},{id:"334285",title:"Ph.D. Student",name:"Sameer",middleName:"Kumar",surname:"Jagirdar",slug:"sameer-jagirdar",fullName:"Sameer Jagirdar",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/334285/images/14691_n.jpg",biography:"I\\'m a graduate student at the center for biosystems science and engineering at the Indian Institute of Science, Bangalore, India. I am interested in studying host-pathogen interactions at the biomaterial interface.",institutionString:null,institution:{name:"Indian Institute of Science Bangalore",country:{name:"India"}}},{id:"329248",title:"Dr.",name:"Md. Faheem",middleName:null,surname:"Haider",slug:"md.-faheem-haider",fullName:"Md. Faheem Haider",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/329248/images/system/329248.jpg",biography:"Dr. Md. Faheem Haider completed his BPharm in 2012 at Integral University, Lucknow, India. In 2014, he completed his MPharm with specialization in Pharmaceutics at Babasaheb Bhimrao Ambedkar University, Lucknow, India. He received his Ph.D. degree from Jamia Hamdard University, New Delhi, India, in 2018. He was selected for the GPAT six times and his best All India Rank was 34. Currently, he is an assistant professor at Integral University. Previously he was an assistant professor at IIMT University, Meerut, India. He has experience teaching DPharm, Pharm.D, BPharm, and MPharm students. He has more than five publications in reputed journals to his credit. Dr. Faheem’s research area is the development and characterization of nanoformulation for the delivery of drugs to various organs.",institutionString:"Integral University",institution:{name:"Integral University",country:{name:"India"}}},{id:"329795",title:"Dr.",name:"Mohd Aftab",middleName:"Aftab",surname:"Siddiqui",slug:"mohd-aftab-siddiqui",fullName:"Mohd Aftab Siddiqui",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/329795/images/15648_n.jpg",biography:"Dr. Mohd Aftab Siddiqui is currently working as Assistant Professor in the Faculty of Pharmacy, Integral University, Lucknow for the last 6 years. He has completed his Doctor in Philosophy (Pharmacology) in 2020 from Integral University, Lucknow. He completed his Bachelor in Pharmacy in 2013 and Master in Pharmacy (Pharmacology) in 2015 from Integral University, Lucknow. He is the gold medalist in Bachelor and Master degree. He qualified GPAT -2013, GPAT -2014, and GPAT 2015. His area of research is Pharmacological screening of herbal drugs/ natural products in liver and cardiac diseases. He has guided many M. Pharm. research projects. He has many national and international publications.",institutionString:"Integral University",institution:null},{id:"333824",title:"Dr.",name:"Ahmad Farouk",middleName:null,surname:"Musa",slug:"ahmad-farouk-musa",fullName:"Ahmad Farouk Musa",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/333824/images/22684_n.jpg",biography:"Dato’ Dr Ahmad Farouk Musa\nMD, MMED (Surgery) (Mal), Fellowship in Cardiothoracic Surgery (Monash Health, Aust), Graduate Certificate in Higher Education (Aust), Academy of Medicine (Mal)\n\n\n\nDato’ Dr Ahmad Farouk Musa obtained his Doctor of Medicine from USM in 1992. He then obtained his Master of Medicine in Surgery from the same university in the year 2000 before subspecialising in Cardiothoracic Surgery at Institut Jantung Negara (IJN), Kuala Lumpur from 2002 until 2005. He then completed his Fellowship in Cardiothoracic Surgery at Monash Health, Melbourne, Australia in 2008. He has served in the Malaysian army as a Medical Officer with the rank of Captain upon completing his Internship before joining USM as a trainee lecturer. He is now serving as an academic and researcher at Monash University Malaysia. He is a life-member of the Malaysian Association of Thoracic & Cardiovascular Surgery (MATCVS) and a committee member of the MATCVS Database. He is also a life-member of the College of Surgeons, Academy of Medicine of Malaysia; a life-member of Malaysian Medical Association (MMA), and a life-member of Islamic Medical Association of Malaysia (IMAM). Recently he was appointed as an Interim Chairperson of Examination & Assessment Subcommittee of the UiTM-IJN Cardiothoracic Surgery Postgraduate Program. As an academic, he has published numerous research papers and book chapters. He has also been appointed to review many scientific manuscripts by established journals such as the British Medical Journal (BMJ). He has presented his research works at numerous local and international conferences such as the European Association for Cardiothoracic Surgery (EACTS) and the European Society of Cardiovascular Surgery (ESCVS), to name a few. He has also won many awards for his research presentations at meetings and conferences like the prestigious International Invention, Innovation & Technology Exhibition (ITEX); Design, Research and Innovation Exhibition, the National Conference on Medical Sciences and the Annual Scientific Meetings of the Malaysian Association for Thoracic and Cardiovascular Surgery. He was awarded the Darjah Setia Pangkuan Negeri (DSPN) by the Governor of Penang in July, 2015.",institutionString:null,institution:{name:"Monash University Malaysia",country:{name:"Malaysia"}}},{id:"30568",title:"Prof.",name:"Madhu",middleName:null,surname:"Khullar",slug:"madhu-khullar",fullName:"Madhu Khullar",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/30568/images/system/30568.jpg",biography:"Dr. Madhu Khullar is a Professor of Experimental Medicine and Biotechnology at the Post Graduate Institute of Medical Education and Research, Chandigarh, India. She completed her Post Doctorate in hypertension research at the Henry Ford Hospital, Detroit, USA in 1985. She is an editor and reviewer of several international journals, and a fellow and member of several cardiovascular research societies. Dr. Khullar has a keen research interest in genetics of hypertension, and is currently studying pharmacogenetics of hypertension.",institutionString:"Post Graduate Institute of Medical Education and Research",institution:{name:"Post Graduate Institute of Medical Education and Research",country:{name:"India"}}},{id:"223233",title:"Prof.",name:"Xianquan",middleName:null,surname:"Zhan",slug:"xianquan-zhan",fullName:"Xianquan Zhan",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/223233/images/system/223233.png",biography:"Xianquan Zhan received his MD and Ph.D. in Preventive Medicine at West China University of Medical Sciences. He received his post-doctoral training in oncology and cancer proteomics at the Central South University, China, and the University of Tennessee Health Science Center (UTHSC), USA. He worked at UTHSC and the Cleveland Clinic in 2001–2012 and achieved the rank of associate professor at UTHSC. Currently, he is a full professor at Central South University and Shandong First Medical University, and an advisor to MS/PhD students and postdoctoral fellows. He is also a fellow of the Royal Society of Medicine and European Association for Predictive Preventive Personalized Medicine (EPMA), a national representative of EPMA, and a member of the American Society of Clinical Oncology (ASCO) and the American Association for the Advancement of Sciences (AAAS). He is also the editor in chief of International Journal of Chronic Diseases & Therapy, an associate editor of EPMA Journal, Frontiers in Endocrinology, and BMC Medical Genomics, and a guest editor of Mass Spectrometry Reviews, Frontiers in Endocrinology, EPMA Journal, and Oxidative Medicine and Cellular Longevity. He has published more than 148 articles, 28 book chapters, 6 books, and 2 US patents in the field of clinical proteomics and biomarkers.",institutionString:"Shandong First Medical University",institution:{name:"Affiliated Hospital of Shandong Academy of Medical Sciences",country:{name:"China"}}},{id:"297507",title:"Dr.",name:"Charles",middleName:"Elias",surname:"Assmann",slug:"charles-assmann",fullName:"Charles Assmann",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/297507/images/system/297507.jpg",biography:"Charles Elias Assmann is a biologist from Federal University of Santa Maria (UFSM, Brazil), who spent some time abroad at the Ludwig-Maximilians-Universität München (LMU, Germany). He has Masters Degree in Biochemistry (UFSM), and is currently a PhD student at Biochemistry at the Department of Biochemistry and Molecular Biology of the UFSM. His areas of expertise include: Biochemistry, Molecular Biology, Enzymology, Genetics and Toxicology. He is currently working on the following subjects: Aluminium toxicity, Neuroinflammation, Oxidative stress and Purinergic system. Since 2011 he has presented more than 80 abstracts in scientific proceedings of national and international meetings. Since 2014, he has published more than 20 peer reviewed papers (including 4 reviews, 3 in Portuguese) and 2 book chapters. He has also been a reviewer of international journals and ad hoc reviewer of scientific committees from Brazilian Universities.",institutionString:"Universidade Federal de Santa Maria",institution:{name:"Universidade Federal de Santa Maria",country:{name:"Brazil"}}},{id:"217850",title:"Dr.",name:"Margarete Dulce",middleName:null,surname:"Bagatini",slug:"margarete-dulce-bagatini",fullName:"Margarete Dulce Bagatini",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/217850/images/system/217850.jpeg",biography:"Dr. Margarete Dulce Bagatini is an associate professor at the Federal University of Fronteira Sul/Brazil. She has a degree in Pharmacy and a PhD in Biological Sciences: Toxicological Biochemistry. She is a member of the UFFS Research Advisory Committee\nand a member of the Biovitta Research Institute. She is currently:\nthe leader of the research group: Biological and Clinical Studies\nin Human Pathologies, professor of postgraduate program in\nBiochemistry at UFSC and postgraduate program in Science and Food Technology at\nUFFS. She has experience in the area of pharmacy and clinical analysis, acting mainly\non the following topics: oxidative stress, the purinergic system and human pathologies, being a reviewer of several international journals and books.",institutionString:"Universidade Federal da Fronteira Sul",institution:{name:"Universidade Federal da Fronteira Sul",country:{name:"Brazil"}}}]}},subseries:{item:{id:"38",type:"subseries",title:"Pollution",keywords:"Human activity, Pollutants, Reduced risks, Population growth, Waste disposal, Remediation, Clean environment",scope:"
\r\n\tPollution is caused by a wide variety of human activities and occurs in diverse forms, for example biological, chemical, et cetera. In recent years, significant efforts have been made to ensure that the environment is clean, that rigorous rules are implemented, and old laws are updated to reduce the risks towards humans and ecosystems. However, rapid industrialization and the need for more cultivable sources or habitable lands, for an increasing population, as well as fewer alternatives for waste disposal, make the pollution control tasks more challenging. Therefore, this topic will focus on assessing and managing environmental pollution. It will cover various subjects, including risk assessment due to the pollution of ecosystems, transport and fate of pollutants, restoration or remediation of polluted matrices, and efforts towards sustainable solutions to minimize environmental pollution.
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Dr. Rahman was also adjunctly attached with Kanazawa University, Japan (Visiting Research Professor, Dec 2014 to Mar 2015; JSPS Postdoctoral Research Fellow, Apr 2012 to Mar 2014), and Tokyo Institute of Technology, Japan (TokyoTech-UNESCO Research Fellow, Oct 2004–Sep 2005). \nHe received his Ph.D. degree in Environmental Analytical Chemistry from Kanazawa University, Japan (2011). He also achieved a Diploma in Environment from the Tokyo Institute of Technology, Japan (2005). Besides, he has an M.Sc. degree in Applied Chemistry and a B.Sc. degree in Chemistry, all from the University of Chittagong, Bangladesh. \nDr. Rahman’s research interest includes the study of the fate and behavior of environmental pollutants in the biosphere; design of low energy and low burden environmental improvement (remediation) technology; implementation of sustainable waste management practices for treatment, handling, reuse, and ultimate residual disposition of solid wastes; nature and type of interactions in organic liquid mixtures for process engineering design applications.",institutionString:null,institution:{name:"Fukushima University",institutionURL:null,country:{name:"Japan"}}},editorTwo:{id:"201020",title:"Dr.",name:"Zinnat Ara",middleName:null,surname:"Begum",slug:"zinnat-ara-begum",fullName:"Zinnat Ara Begum",profilePictureURL:"https://mts.intechopen.com/storage/users/201020/images/system/201020.jpeg",biography:"Zinnat A. 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